Early Life History and Growth Chapters 9 & 10 2 growth traits set fish apart from other vertebrates: Indeterminate Growth continual increase in length and volume Larval Stage larval stage usually bears little resemblance to juvenile or adult Adult Larval- Juvenile 1
Early Life History - Gametogenesis 1) Gametogenesis - Spermatogenesis (sperm very variable) - Oogenesis (oocytes w/ yolk) - Vitellogenesis yolk granules Early Life History - Fecundity 2) Fecundity = number of eggs released by a female 1 or 2 up to millions; depends on life history v. Fertility (fish reabsorb unused eggs); 2
Early Life History - Fecundity 2) Fertility v. Fecundity (based on egg counts) Most marine fishes = pelagic and external fertilization; Most Freshwater = parental care/bottom or vegetation or nests. Demersal laid on bottom Early Life History Reproductive Effort 3) Reproductive Effort and Activity Look at eggs instantaneous versus cumulative measures Instantaneous vs. Cumulative Gonadosomatic index GSI usually % weight 5% (cichlids), salmonids (20%- 30%) to 47% (eels) males much lower 3
Early Life History Reproductive Effort 3) Gonadosomatic index Good for total spawners Underestimates repeat, batch or serial spawners Early Life History - Fertilization 4) Fertilization external in most; internal in all sharks and a few bony families; Some poeciliid live bearers = gynogenetic (use other males sperm to activate); Internal requires males have intromittent organ (claspers, gonopodium) 4
Early Life History Embryology 5) Embryology after fertilization Chorion hardens = water hardening Early Life History Embryology 5) Embryology Oviparous egg laying Viviparous develop inside mother = live bearing about half (+ 500) Chondricthyes; 500 (2%) bony Ovoviviparity inside mother but depend on yolk Development depends on temperature etc causes meristic variation usually colder means more scales, fin rays and vertebrae 5
Early Life History Embryology 5) Embryology Meristic Variation - Development depends on temperature etc causes meristic variation Jordan s rule latitude effects on meristic numbers - usually colder means more scales, fin rays and vertebrae opposite guppies and plaice V relationship fewer at intermediate Early Life History 6) Larvae Free embryo (free swimming young with yolk sac - alevin) to Fry (planktonic food) 6
6) Larvae Early Life History Direct v. indirect development larval stage brief or non-definable ( miniature ) versus distinct metamorphosis 6) Larvae Early Life History Larval Feeding and Survival Food very important right after yolk stage important for population dynamics of commercial species most larvae die in first week from starvation/predation Critical Period Hypothesis Point of No Return 7
Early Life History 6) Larvae Larval Feeding and Survival Match-Mismatch Hypothesis Early Life History 6) Larvae Growth More later under bioenergetics Gross Growth Efficiency - weight increase to weight food Assimilation Efficiency food actually used 8
Early Life History 6) Larvae Larvae usually not like adult spines, large fins etc for anti-predator protection as fish get older (larger) usually less risk of predation. Ecological, physiological and behavioral competence all improve. Movement of larvae interesting/debated many marine fish spawn off shore. Larvae move inshore to weed beds/estuaries etc. wind driven currents, tides etc. Individuals: Life Histories & Growth Chapter 10 Juveniles to Adults - Growth 1) Juvenile when larval features lost vs. miniature adult Growth = metamorphosis Growth = change is size (body material) 9
Life History Characters Age and size at maturation - Early v late = trade-off. More eggs older but may die first; younger fewer eggs and reduced growth and weaker state if they decide to reproduce. Fish under heavy predation reproduce sooner. Growth and Age Growth = any change in size or amount of body material, regardless of whether that change is positive or negative or temporary or long lasting Growth (energetically) = change in calories stored as somatic or gonad tissue (will discuss later) 10
Metamorphosis Lampreys ammocoetes to adult. Asymmetric flatfish. I. Depression (dorsal ventral flat) v. Compression (lateral flat). Ii. Incomplete ossification and rotation. Iii. Left eye flounders - sinistral. Iv. Right eye flounders - dextral. Metamorphosis Smoltification salmonids (fresh to saltwater). Redds eggs alevins (egg sac) fry parr w/ parr marks after months/years go downstream (smolts). Countershade silvery; Streamline loss lipids, more buoyant (inc gas volume) hemoglobins change, gill structure (inc chloride cells) reverse rheotaxis imprint odor. Hormonal control most changes; If don t get to sea they revert to parr and mature quickly (1 yr). 11
Growth and Age - For Time T1 to T2 With Sizes Y1 and Y2 1) Absolute Growth = Y2 Y1 2) Absolute Growth Rate = (Y2 Y1) / (T2 T1) 3) Relative Growth = (Y2 Y1) / Y1 4) Relative Growth Rate = (Y2 Y1) / [Y1 (T2 T1)] Linear process but if exponential use instantaneous growth rates; G = (log e Y2 log e Y1) / (T2 T1) 12
Length Versus Age = Growth Curves Fit model to curve von Bertalanffy growth equation Gompertz equation Lt = Lmax (1 e gt ) T = time units (tx t0), Lt = length at time t, Lmax = maximum length, g = growth coefficient, e = base natural log Model Equations 13
Quantification of Size Changes (Growth) 1) Body length standard length (SL), fork length (FL), total length (TL) Quantification of Size Changes (Growth) Weight traditional measure growth & production Mass & Length Easy to do W = al b where b usually 2.5 to 3.0 W = log a + b log L Good = more accurate as fish get larger; > or < 3 means positive or negative allometric growth Bad = transient, water, lipids, gonads, stomach 14
Other Quantification size changes K = condition factor for fish K = W/L 3 Some use Wet v Dry weight v Ash Weight Other Quantification of size changes Proximate Analysis categories of compounds in a mixture 1) Carbohydrates (low not typical) 2) Proteins w/ M.W. over 10,00 3) Lipids = good indicator 15
Scale and Allometry Scale and Allometry as fish grow dimensions of the body change nonlinear Types of Change: Dimension change (add mass length) Material change (cart to bone) Design change (anguilliform to carangiform swim) Estimations of Growth in Natural Populations Remember most techniques developed in temperate populations 16
Estimate Growth in Natural Populations Length Frequency Distribution in identifiable cohorts Peterson Method. Assume cohorts cluster around mean Advantages easy to collect data; Low tech, anyone can do, need ruler or measuring board. Disadvantage - Hard to analyze data, Bad in continuously breeding populations (tropics), better for young. Estimations Growth in Nat Populations Back calculation Bony parts carry record of growth; otoliths, vertebrae, fin spines, scales, other hard body parts Advantages Good data Disadvantages expensive, time consuming, caution in ring interpretation, tropical no annual marks, need all size and age classes 17
Estimate Growth in Natural Populations Mark Recapture recovering marked fish of known age Tagging - clip fins, paint, tetracycline, pit tags, disks, flags, magnetic implants +++ Good data --- Expensive, time consuming esp in large systems w/poor returns, tags influence, difficult recapture Estimate Growth in Natural Populations Raise in controlled environment Growth Rate = 100 (log e W f -log e W i )/(t f -t i ) Good for aquaculture 18
Growth Usually annual Daily in small fish a) Frasier-Lee L = a + bs b) Regression c) Covariance d) Linear model approach Bioenergetics Model Consumption = Metabolism (Respiration) + Wastes (Excreted) + Growth (Production); C = M + E + G 19
C = M + E + G Bioenergetics Model C = (M = respiration + active metabolism + SDA specific dynamic action) + (E = egestion + excretion) + (G = somatic growth + gonad production) = (R + A + S) + (F + U) + (B + G) Bioenergetics Model C = M + E + G Mass Balance Approach Environmental Stress Growth down if C decreases or M increases 20
Bioenergetics Model C = M + E + G Consumption = proportion of maximum daily ration for fish at particular mass and temperature = maximum rate = g of prey per g body mass per day Bioenergetics Model C = M + E + G Temperature dependence of consumption Different functions 21
Bioenergetics Model C = M + E + G Respiration is also dependent upon fish size, temperature and activity Different functions Bioenergetics Model C = M + E + G Egestion and Excretion is constant proportion of consumption or as function of temperature and consumption Different functions 22
Bioenergetics Model Scaling Individuals to Populations Cohort = group of similar sized (aged) fish of the same species experiencing the same environmental conditions (temperature, diet, growth and reproductive losses) Combine cohorts for population Bioenergetics Model Population Mortality Mortality = important population process Natural and fishing 23